Among-species variation in six decades of changing migration timings explained through ecology, life-history and local migratory abundance.

Species exploiting seasonal environments must alter timings of key life-history events in response to large-scale climatic changes in order to maintain trophic synchrony with required resources. Yet, substantial among-species variation in long-term phenological changes has been observed. Advancing from simply describing such variation towards predicting future phenological responses requires studies that rigorously quantify and explain variation in the direction and magnitude of changing timings across diverse species in relation to key ecological and life-history variables. Accordingly, we fitted multi-quantile regressions to 59 years of multi-species data on spring and autumn bird migration timings through northern Scotland. We demonstrate substantial variation in changes in timings among 72 species, and tested whether such variation can be explained by species ecology, life-history and changes in local abundance. Consistent with predictions, species that advanced their migration timing in one or both seasons had more seasonally restricted diet types, fewer suitable breeding habitat types, shorter generation lengths and capability to produce multiple offspring broods per year. In contrast, species with less seasonally restricted diet types and that produce single annual offspring broods, showed no change. Meanwhile, contrary to prediction, long-distance and short-distance migrants advanced migration timings similarly. Changes in migration timing also varied with changes in local migratory abundance, such that species with increasing seasonal abundance apparently altered their migration timing, whilst species with decreasing abundance did not. Such patterns broadly concur with expectation given adaptive changes in migration timing. However, we demonstrate that similar patterns can be generated by numerical sampling given changing local abundances. Any apparent phenology-abundance relationships should, therefore, be carefully validated and interpreted. Overall, our results show that migrant bird species with differing ecologies and life-histories showed systematically differing phenological changes over six decades contextualised by large-scale environmental changes, potentially facilitating future predictions and altering temporal dynamics of seasonal species co-occurrences.

Quantifying full phenological event distributions reveals simultaneous advances, temporal stability and delays in spring and autumn migration timing in long-distance migratory birds.

Phenological changes in key seasonally expressed life-history traits occurring across periods of climatic and environmental change can cause temporal mismatches between interacting species, and thereby impact population and community dynamics. However, studies quantifying long-term phenological changes have commonly only measured variation occurring in spring, measured as the first or mean dates on which focal traits or events were observed. Few studies have considered seasonally paired events spanning spring and autumn or tested the key assumption that single convenient metrics accurately capture entire event distributions. We used 60 years (1955-2014) of daily bird migration census data from Fair Isle, Scotland, to comprehensively quantify the degree to which the full distributions of spring and autumn migration timing of 13 species of long-distance migratory bird changed across a period of substantial climatic and environmental change. In most species, mean spring and autumn migration dates changed little. However, the early migration phase (≤10th percentile date) commonly got earlier, while the late migration phase (≥90th percentile date) commonly got later. Consequently, species' total migration durations typically lengthened across years. Spring and autumn migration phenologies were not consistently correlated within or between years within species and hence were not tightly coupled. Furthermore, different metrics quantifying different aspects of migration phenology within seasons were not strongly cross-correlated, meaning that no single metric adequately described the full pattern of phenological change. These analyses therefore reveal complex patterns of simultaneous advancement, temporal stability and delay in spring and autumn migration phenologies, altering species' life-history structures. Additionally, they demonstrate that this complexity is only revealed if multiple metrics encompassing entire seasonal event distributions, rather than single metrics, are used to quantify phenological change. Existing evidence of long-term phenological changes detected using only one or two metrics should consequently be interpreted cautiously because divergent changes occurring simultaneously could potentially have remained undetected.

Decline in an Atlantic Puffin Population: Evaluation of Magnitude and Mechanisms.

Determining which demographic and ecological parameters contribute to variation in population growth rate is crucial to understanding the dynamics of declining populations. This study aimed to evaluate the magnitude and mechanisms of an apparent major decline in an Atlantic Puffin Fratercula arctica population. This was achieved using a 27-year dataset to estimate changes in population size and in two key demographic rates: adult survival and breeding success. Estimated demographic variation was then related to two ecological factors hypothesised to be key drivers of demographic change, namely the abundance of the main predator at the study site, the Great Skua Stercorarius skua, and Atlantic Puffin chick food supply, over the same 27-year period. Using a population model, we assessed whether estimated variation in adult survival and reproductive success was sufficient to explain the population change observed. Estimates of Atlantic Puffin population size decreased considerably during the study period, approximately halving, whereas Great Skua population estimates increased, approximately trebling. Estimated adult Atlantic Puffin survival remained high across all years and did not vary with Great Skua abundance; however, Atlantic Puffin breeding success and quantities of fish prey brought ashore by adults both decreased substantially through the period. A population model combining best possible demographic parameter estimates predicted rapid population growth, at odds with the long-term decrease observed. To simulate the observed decrease, population models had to incorporate low immature survival, high immature emigration, or increasingly high adult non-breeding rates. We concluded that reduced recruitment of immatures into the breeding population was the most likely cause of population decrease. This study showed that increase in the size of a predator population does not always impact on the survival of adult prey and that reduced recruitment can be a crucial determinant of seabird population size but can easily go undetected.